1. Ecology: Animal Behavior
The study of behavior is known as ethology.

2. Responding to a Changing Environment
1. Physiological Responses - changing the functioning of the body - acclimation (dilating capillaries to release heat) 2. Morphological Responses - changing the anatomy (structure) of the body - growing thicker fur or change in fur color in winter 3. Behavioral Responses - changing behavior to adapt to the change - moving to a more favorable location - cooperative behavior - agonistic behavior when threatened
Ask students to come up with additional examples of how animals adapt to changing conditions.

3. Nature vs. Nurture? Genes and the environment both influence behavior
Innate behavior is developmentally fixed, regardless of the environment, and under strong genetic influence (ex. agonistic behavior)
Learned behavior is due to cognitive development, change with experience & environment (ex. Mother bears teach their cubs about hunting, berry picking, fishing, and the best places to find tasty grubs) 
A behavior is the nervous system’s response to a stimulus and is carried out by the muscular or the hormonal system. Behavior is subject to natural selection. “”Nature” refers to the animal’s genetics while “nurture” refers to the animal’s environment, especially with regard to learned behaviors. The debate between the two has been ongoing in biology.
In 1979, Thomas Bouchard began to study twins who were separated at birth and reared in different families. He found that an identical twin reared away from his or her co-twin seems to have about an equal chance of being similar to the co-twin in terms of personality, interests, and attitudes as one who has been reared with his or her co-twin. This leads to the conclusion that the similarities between twins are due to genes, not environment, since the differences between twins reared apart must be due totally to the environment.

4. Proximate and Ultimate Questions
Proximate questions focus on the environmental stimuli that trigger behavior
Physiological & genetic mechanisms of behavior
How does a behavior happen ?
Ultimate questions focus on the evolutionary significance of a behavior
Why does a behavior happen?
What is the evolutionary benefit of the behavior?
Behavioral ecology integrates proximate and ultimate explanations for animal behavior
Proximate causation addresses “how” a behavior occurs or is modified
Ultimate causation addresses “why” a behavior occurs in the context of natural selection
Natural selection plays a huge role in the evolution of these behaviors:
a. Animal behavior ties directly to differing reproductive success rates. (Beneficial behavior vs. Harmful Behavior)
b. Animals possessing beneficial or “strong” traits have a higher survival rate, thus produce more offspring possessing “strong” traits.
c. Animals possessing “weaker” traits often die before reaching reproductive age, thus weeding out the genes that were “weak” or less beneficial.

5. Competitive Social Behaviors Often Represent Contests for Resources
Cooperative behavior is when an animal invests resources in a common interest shared by other group members
Agonistic behavior is made of a suite of three different divisions of behaviors: threats, aggression, and submission.

6. Competitive Social Behaviors Often Represent Contests for Resources
Agonistic behavior is any social behavior that involves fighting, thus it is a contest involving threats.
Agonistic behavior is made of a suite of three different divisions of behaviors: threats, aggression, and submission.
Generally, no harm is done

7. Competitive Social Behaviors Often Represent Contests for Resources
Reconciliation behavior often happens between conflicting individuals.
Many animals “kiss and make up”.

8. Competitive Social Behaviors Often Represent Contests for Resources
Dominance hierarchies involve a ranking of individuals in a group (a “pecking order”).
Alpha and beta rankings exist, the alpha organisms control the behavior of others.
In herd animals the alpha often does extra work.

9. Fixed Action Patterns (FAP)
FAP is a sequence of unlearned (innate) , unchangeable behavioral acts, that once started, are carried out to completion
Triggered by a sign stimulus (external sensory stimulus)
Ex: Agonistic (aggressive) behavior in 3-spined stickleback male fish in response to the red underside of an intruder fish
LO 3.42 The student is able to describe how organisms exchange information in response to internal changes or environmental cues.
Fixed Action Patterns
A fixed action pattern is a sequence of unlearned, innate behaviors that is unchangeable. Once initiated, it is usually carried to completion
A fixed action pattern is triggered by an external cue known as a sign stimulus. Selection for individual survival and reproductive success can explain most behaviors. Behavior enhances survival and reproductive success in a population.

10. A Recreation of Tinbergen’s Experiment with Three-Spined Stickleback and Fixed Action Patterns
Niko Tinbergen was a pioneer in the study of animal behavior. He suggested that understanding any behavior requires answering FOUR questions:
What stimulus elicits the behavior, and what physiological mechanisms mediate the response?
How does the animal’s experience during growth and development influence the response?
How does the behavior aid survival and reproduction?
What is the behavior’s evolutionary history?

11. Imprinting
Imprinting is a type of behavior that includes both learning and innate components and is irreversible
limited phase early in an animal’s development, is the only time certain behaviors can be learned (critical period)
Incubator-hatched goslings imprinted on scientist (Konrad Lorenz) during first few hours of life and followed him

12. Ever Seen the Movie “Fly Away Home”?
The movie involves Canadian Geese, but the concept is based on an organization named Operation Migration which has played a leading role in the reintroduction of endangered Whooping cranes into eastern North America since In the 1940s the species was reduced to just 15 birds.

13. Directed Movements Controlled by genes
Kinesis = change in activity rate in response to stimuli. Ex. Isopods live best in moist conditions; move more in dry areas to increases likelihood of encountering a moist area
Taxis = a more or less automatic, oriented movement toward or away from a stimulus
Migration - using sun (seasonal changes), stars, Earth’s magnetic field, etc.
Mnemonic Device: Kinetic refers to movement as in kinetic energy so it is easy to remember Kinesis is about random movement in response to a change in a stimulus, such as increased movement when it is warmer, but not toward or away from the heat source.
Taxis – you take a taxi to a specific location for a reason – Taxis is movement “for a reason” i.e. in response to a stimulus one moves either towards or away from the stimulus – they “take a taxi to get there”.

14. Animal Signals & Communication
Signal - behavior that causes change in another’s behavior
Communication involves the transmission of, reception of, and response to signals between animals
Animal communication is any behavior on the part of one animal that has an effect on the current or future behavior of another animal.

15. Animal Signals & Communication
Chemical Communication:
Pheromones – important in reproduction behavior
Scents—important in marking territory or defense
Auditory Communication (vocalization):
Drosophila males produce a characteristic “song” by beating their wings, insects (innate, genetic)
Mating songs in birds (innate & learned)
Many mammals, in particular, have glands that generate distinctive and long-lasting smells, and have corresponding behaviors that leave these smells in places where they have been. Often the scented substance is introduced into urine or feces. Bees carry with them a pouch of material from the hive which they release as they reenter, the smell of which indicates that they are a part of the hive and grants their safe entry. Ants use pheromones to create scent trails to food as well as for alarm calls, mate attraction and to distinguish between colonies. Additionally, they have pheromones that are used to confuse an enemy and manipulate them into fighting with each other.
Male gray tree frogs (Hyla versicolor) give out longer but fewer calls in reaction to the calls of other males. In other words, when these frogs are chorusing full blast, a male seeking female attention will change the rhythm of his call to break out of the chorus. 

16. Territoriality
Territorial animals defend areas that contain a nest, den or mating site and sufficient food resources for themselves and their young.
Territoriality uses a great deal of an individual’s energy
In addition, an individual might die defending a territory, thus miss a reproductive opportunity
Spraying behavior is a way for an individual to mark its territory
Ask student for example of animals that exhibit this behavior. The red breast of the robin (visual), bird songs or the calls of gibbons (auditory), or through the deposit of scent marks (olfactory). Bears, dogs, cats, lemurs, etc. use scent-marking to signal the boundaries of their territories.

17. Environment & Genetics
Environmental factors, such as the quality of the diet, the nature of social interactions, and opportunities for learning can influence the development of behaviors in every group of animals
Example: Variations in diet led to rejection of mates in Drosophila
How is it connected to genetics? It relates to changes in gene expression.
Diane Dodd raised two strains of fruit flies on different diets (one on starch and one on maltose) after 25 generations, the menu affected mate selection. The change in diet was an artificial evolutionary pressure that led to a evolutionary change in mating habits which occurs via gene expression.

18. Learning
Learning is the modification of behavior based on specific experiences
Maturation: behavior due to developing physiological changes
Habituation: loss of responsiveness to stimuli that convey little or no information
Spatial Learning: the modification of behavior based on experience with the spatial structure of the environment, including the location of nest sites, hazards, food, and prospective mates
Associative Learning- behavior through trial and error
Ask students to provide examples of each of these types of learning.
Maturation: Human babies learning to sit up, crawl, walk, run, etc. vs. calves that stand up and walk within minutes of being born
Habituation: “crying wolf”, parents nagging, odors!! (olfactory fatigue)
Spatial Learning: A young human being able to judge when it is safe to cross the street and eventually walk home from school without getting lost
Associative Learning-A young human learning their addition, multiplication facts or mastering any repetitive task such as making a basket in basketball, etc.

19. Associative Learning
The ability of many animals to associate one feature of the environment with another
Classical Conditioning
arbitrary stimulus associated with reward or punishment
Pavlov’s Experiment
Operant Conditioning
“trial-and-error learning”
Associates behavior with reward or punishment, Skinner Box
LO 2.40The student is able to connect concepts in and across domain(s) to predict how environmental factors affect responses to information and change behavior.
Bird Flocking
Simulator:
Dolphins & Bats Biosonar
EvoTutor
Pavlov

20. Cognition & Problem Solving
The ability of animal’s nervous system to perceive, store, process, and use info gathered by sensory receptors
Ex: monkey stacking boxes in order to reach bananas or the use of tools

21. Natural Selection & Behaviorism
Genetic components of behavior evolve through natural selection which favors behaviors that increase survival and reproductive success
Foraging behavior – Balance between benefits of nutrition and cost of finding food (predation, energy, etc.)
Cost-benefit analysis
Mate selection
Most animals are promiscuous – no strong pair bonds
Monogamous – one male with only one female
Polygamous – an individual of one sex mating with several of the other
Polygyny – one male with many females
Polyandry – one female with many males
Anytime an organism can reserve energy to reproduce they are increasing their fitness and the chances that their genes will be passed on through their offspring. Foraging behavior increases fitness.
Reproductive fitness and Energy--Those organisms that save energy on foraging will mostly use that saved energy to reproduce, thus the most fit genes are passed on to offspring with frequency.

22. Interpret This Data
Optimal foraging theory (a cost analysis of energy input vs. energy yield.) Ask students to interpret the data given knowing that crows drop the whelks (small sea snails) onto the rocks of the islands located off British Columbia in order to crack their shells so they can eat the soft insides. If the shell doesn’t break, the crow tries and tries again. Higher flight = greater acceleration of falling whelk = increased impact of whelk on rocks = increased chance of destroying shell (simple physics!)
Crows that learn to drop the whelk from 5+ m get the most energy out of their food vs. energy input (expended as a result of having to fly higher). Those that fly below 5+ m learn they will have to expend MORE energy making additional flights in order to crack the whelk. Those that fly higher than 5+ m are also wasting more energy than need be, so it works both ways!

23. (Drop Height  Navg Drops) = “Cost−Benefit Analysis”
Crow & Whelk Data
Navg Drops Required
to Break Shell
(Drop Height  Navg Drops) = “Cost−Benefit Analysis”
Drop Height (m)
5 6 30
7 5 35
Again, ask students to interpret the data presented. Point out that Navg symbolizes the “Average number of drops”. Also have them interpret the significance of the final column. The lowest number in that column represents the most efficient use of energy expended to accomplish the task!
Foraging is a food-obtaining behavior. The optimal foraging theory states that natural selection will benefit animals that maximize their energy intake-to-expenditure ratio, thus flying at 5 m results in the most energy efficient process.
Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

24. The Concept Of Inclusive Fitness Can Account For Most Altruistic Behavior
Most social behaviors are selfish, so how do we account for behaviors that help others?
Altruism is defined as behavior that might decrease individual fitness, but increase the fitness of others.
Ask students for examples of altruistic behaviors and animals that exhibit them.
Vampire bats regularly regurgitate blood and donate it to other members of their group who have failed to feed that night, ensuring they do not starve.
In numerous bird species, a breeding pair receives help in raising its young from other ‘helper’ birds, who protect the nest from predators and help to feed the fledglings.
Vervet monkeys give alarm calls to warn fellow monkeys of the presence of predators, even though in doing so they attract attention to themselves, increasing their personal chance of being attacked.

25. Altruism
Altruistic behavior is common throughout the animal kingdom, particularly in species with complex social structures.
In social insect colonies (ants, wasps, bees and termites), sterile workers devote their whole lives to caring for the queen, constructing and protecting the nest, foraging for food, and tending the larvae.
Such behavior is maximally altruistic: sterile workers obviously do not leave any offspring of their own — so have personal fitness of zero — but their actions greatly assist the reproductive efforts of the queen.
Altruism is a great puzzle for evolution to explain – ask students to offer possible answers as to how altruism might evolve, then go on to discuss kin selection, etc., though those are not the only possible answers, but they are the ones most likely to be on the exam!

26. Inclusive Fitness
The inclusive fitness of an organism is the sum of its classical fitness (how many of its own offspring it produces and supports) and the number of equivalents of its own offspring it can add to the population by supporting others.
Advocates of inclusive fitness theory say that an organism can improve its overall genetic success by cooperative social behavior.

27. Inclusive Fitness
From the gene's point of view, evolutionary success ultimately depends on leaving behind the maximum number of copies of itself in the population which would translate into a maximum number of offspring.
In 1964 W. D. Hamilton proved mathematically that, because close relatives of an organism share some identical genes, a gene can also increase its evolutionary success by promoting the reproduction and survival of these related or otherwise similar individuals

28. Kin Selection
Kin selection refers to natural selection that favors altruistic behavior by enhancing reproductive success of relatives.
The graph above shows that female ground squirrels tend to stay close to their relatives whereas male ground squirrels tend to move farther away from their relatives. Females have a higher incidence of altruistic behavior since it increases their fitness in a group when an alarm is given in response to a threat.
Kin selection is the mechanism of inclusive fitness, where individuals help relatives raise young.
Reciprocal altruism, where an individual aids other unrelated individuals without any benefit, is rare, but sometimes seen in primates (often in humans).

29. Kin Selection The basic idea of kin selection is simple.
Imagine a gene which causes its bearer to behave altruistically towards other organisms, e.g. by sharing food with them.
Organisms without the gene are selfish — they keep all their food for themselves, and sometimes get handouts from the altruists.
Clearly the altruists will be at a fitness disadvantage, so we should expect the altruistic gene to be eliminated from the population.
However, suppose that altruists are discriminating in who they share food with. They do not share with just anybody, but only with their relatives..

30. Kin Selection
So when an organism carrying the altruistic gene shares his food, there is a certain probability that the recipients of the food will also carry copies of that gene.
This means that the altruistic gene can in principle spread by natural selection.
The gene causes an organism to behave in a way which reduces its own fitness but boosts the fitness of its relatives — who have a greater than average chance of carrying the gene themselves.
So the overall effect of the behavior may be to increase the number of copies of the altruistic gene found in the next generation, and thus the incidence of the altruistic behavior itself.